Nervous System Nervous System The nervous system is divided into two major parts, (1) the central nervous system (CNS) and (2) the peripheral nervous system (PNS). • Central Nervous System – The central nervous system consists of the brain and the spinal cord. Functions of the central nervous system include integration, control, consciousness, and mental activity. • Peripheral Nervous System – The peripheral nervous system consists of all the nervous system components, such as the nerves and neurons, that extend from or are located outside of the central nervous system. The peripheral nervous system is divided into the (1) sensory (afferent) division and the (2) motor (efferent) division – Afferent (sensory) division • The sensory division involves information collected from the somatic division, visceral division, and special senses and delivered to the central nervous system (CNS). – Efferent (motor) division Anatomy and Physiology Text and Laboratory Workbook, Stephen G. Davenport, Copyright 2006, All Rights Reserved, no part of this publication can be used for any commercial purpose. Permission requests should be addressed to Stephen G. Davenport, Link Publishing, P.O. Box 15562, San Antonio, TX, 78212 • The efferent division involves information flow from the central nervous system (CNS) to the somatic division and the visceral division. Nervous System • Somatic Division – The somatic division of the peripheral nervous system is the division involved with the voluntary control of body movements. The somatic division is divided into the sensory (afferent) and the motor (efferent) components. The sensory component functions in receiving stimuli and conducting information to the CNS concerning voluntary body movements. The motor (efferent) component functions to deliver information from the CNS to the skeletal muscles, thus, directing their contraction. • Visceral Division – The visceral division of the peripheral nervous system is the division involved with involuntary control of body movements such as those of the cardiovascular, digestive, urinary, respiratory, etc., systems. The sensory component functions in receiving stimuli and conducting information concerning involuntary control to the CNS. The motor component functions to deliver information from the CNS to control involuntary movements. The motor component for involuntary control is routed through the autonomic nervous system (ANS). The autonomic nervous system, divided into the parasympathetic and sympathetic divisions, directs motor control to smooth muscle, cardiac muscle, and glands. • Special Senses – The special senses include taste, hearing, equilibrium, vision, and smell. Information from the five senses is routed to the CNS by way of the afferent (sensory) division. Information processed by the central nervous system may produce an efferent (motor) response that is directed to either/both the efferent somatic or visceral divisions. Figure 17.1 An overview of the organization of the nervous system. Neurons • Neurons (nerve cells) are the cells of the nervous system that function in NEURONS – (1) the generation and – (2) the conduction of the nerve impulse, and the – (3) secretion of a neurotransmitter at their terminals. • Neurons have a cell body with one or more processes (nerve fibers) extending from them. 1 Structure of a Multipolar Neuron – Cell Body • Neuron Cell Body – The cell bodies of neurons are located in the gray matter of the central nervous system and in structures called ganglia in the peripheral nervous system. The cell body of a neuron contains abundant cytoplasm with numerous organelles. The organelles that are easily observed with the light microscope include • (1) a large nucleus that contains one to several dark-stained nucleoli and • (2) dark-stained granules called Nissl substance, or bodies (rough endoplasmic reticulum). Structure of a Multipolar Neuron - Processes • Dendrites – Depending upon the type of neuron, one to many processes called dendrites may be present at the cell body. – Dendrites have traditionally been described as the neuron’s processes that function in the conduction of impulses toward its body. – Another description is that the dendrites are the structures that function as the receptive portions of the neuron. The latter function is the one used is this study. Figure 17.2 A “typical” multipolar neuron showing numerous processes associated with the cell body. Only one axon originates at the cell body, all the other processes are dendrites. Structure of a Multipolar Neuron - Processes • Axon – Usually, a neuron has only one process called an axon associated with its cell body. – The site on the cell body where the axon originates is called the axon hillock. – Once the axon leaves the cell body, the axon can split into branches called collaterals. – The axon ends in fine branches called telodendria. The end of each tenodendrion is called an axon terminal that functions in a synapse, the site of where the nerve impulse passes to another neuron, a muscle, or a gland. – Traditionally, an axon has been described as the portion of the neuron that functions in the conduction of impulses away from the cell’s body. – Another description is that the axon is the process that • generates and • conducts the impulse, and • releases a neurotransmitter at its axon terminals. Classification According to Function CLASSIFICATION OF NEURONS Classification According to Function Three classifications of neurons according to function are (1) sensory neurons, (2) motor neurons, and (3) association neurons (interneurons). • Sensory neurons – Neurons that transmit impulses generated at their receptors toward the central nervous system are sensory, or afferent, neurons. They constitute the sensory (afferent) division of the peripheral nervous system. • Motor neurons – Neurons that transmit impulses from the central nervous system to effectors (glands and muscles) are motor, or efferent, neurons. They constitute the motor (efferent) division of the peripheral nervous system. • Association neurons (Interneurons) – Neurons of the CNS that transmit impulses from one neuron to another are generally called association neurons, or interneurons. 2 Classification According to Structure Three classifications of neurons according to structure are (1) unipolar neurons, (2) bipolar neurons, and (3) multipolar neurons. • Unipolar neurons – A unipolar neuron has a single continuous fibrous process that is associated with its body. Its single process is produced by the merging of its receptive dendrites with the conductive axon that terminates with synaptic contacts either in the brain or spinal cord. Unipolar neurons are sensory neurons associated with the peripheral nervous system. • Bipolar neurons – A bipolar neuron has two fibrous process, each process is associated with the cell’s body. Bipolar neurons are sensory neurons associated with the neural pathways of the senses involving sight, hearing, and smell. • Figure 17.3 An overview of the functional classification of neurons. Multipolar neurons – A multipolar neuron has more than two processes associated with its cell body. Only one of the processes is the axon, all of the other processes are dendrites. Multipolar neurons are common in the CNS functioning as association (interneurons), and exiting the CNS functioning as motor neurons. NEUROGLIA Neuroglia are the cells and their associated branching fibers that support neural tissue Figure 17.4 An overview of the structural classification of neurons. Neuroglia Neuroglia are the cells and their associated branching fibers that support neural tissue. • Central nervous system – Four varieties of neuroglia found in the are • • • • (1) ependymal cells, (2) astrocytes, (3) oligodendrocytes, and (4) microglia. • Peripheral Nervous System – Two varieties of neuroglia found in the peripheral nervous system are • (1) satellite cells and • (2) Schwann cells. Neuroglia of the Central Nervous System • Ependymal cells – Ependymal cells are found lining the cerebrospinal fluid containing cavities of the CNS. In the brain ependymal cells line the ventricles, and in the spinal cord they line the central canal. The ependymal cells function in the production and regulation of cerebrospinal fluid (CSF). • Astrocytes – Astrocytes, the most numerous glial cells, are glial cells named for their star-shape. Among their functions are support and nutrient exchange/regulation between neurons and adjacent capillaries. 3 Neuroglia of the Central Nervous System • Oligodendrocytes – Oligodendrocytes are the glial cells that produce the myelin sheaths of CNS axons. The oligodendroglia produces sheet-like extensions that form the myelin sheets. • Microglia – Microglia are phagocytic cells of the CNS. They remove debris, waste, pathogens, and other materials. Neuroglia of the Peripheral Nervous System • Satellite cells – Satellite cells are the glial cells that surround the cell bodies of neurons in ganglia, the only sites in the PNS that contain cell bodies of neurons. • Schwann cells – Schwann cells are the glial cells that are associated with all axons in the PNS. Schwann cells either tightly wrap axons to produce myelin sheaths (myelinated axons) or remain in close association to produce unmyelinated axons. Lab Activity 1 Motor Nerve Cells • Observe a microscopic preparation of “Motor nerve cells, smear” (Nerve cells, spinal cord smear). – Identify the multipolar neurons and neuroglia cells (nuclei). – Locate an isolated neuron and identify its cell body, nucleus, Nissl substance (bodies), and processes. Figure 17.5 Scanning power photograph of a multipolar neuron from a slide preparation labeled “Motor nerve cells, spinal cord smear.” Lab Activity 1 Motor Nerve Cells • Cell body – The cell body contains most of the cell’s organelles and cytoplasm. The cytoplasm contains the usual cell organelles except centrioles (the lack of centrioles makes the cells amitotic). The nucleus is easily observed with one to several nucleoli. Dark-stained areas of rough endoplasmic reticulum called Nissl substance (bodies) can usually be observed with high magnification. Lab Activity 1 Motor Nerve Cells • Neuroglia – The supporting cells of the nerve tissue, the neuroglia, are seen as the dark-stained nuclei distributed throughout the preparation. Mostly consisting of astrocytes, the neuroglia are highly traumatized during tissue preparation leaving their nuclei scattered throughout the preparation. • Cell processes – Usually, on smear preparations the cells are severely traumatized, which makes the microscopic identification of the dendrites and the single axon difficult. The axon is long, may show distal branches, and does not contain Nissl substance (bodies). In comparison, the dendrites are short, have numerous branches, and contain Nissl substance. 4 Figure 17.7 High power photograph of multipolar nerve cells from a slide preparation labeled “Motor nerve cells, spinal cord smear.” Figure 17.6 Low power photograph of motor nerve cells from a slide preparation labeled “Motor nerve cells, spinal cord smear.” The general structure of multipolar neurons is observed. Myelinated and Unmyelinated Axons of the PNS MYELINATED and UNMYELINATED AXONS Peripheral Nervous System Figure 17.8 Illustration showing structural differences between myelinated and unmyelinated axons in the peripheral nervous system. • In the peripheral nervous system, neuroglia called Schwann cells are arranged sequentially along all axons. The plasma membranes of Schwann cells contain a phospholipid called myelin. – If the Schwann cells surround and tightly wrap an axon, they produce a myelinated axon with each Schwann cell producing an area of concentrically wrapped plasma membrane called the myelin sheath. The Schwann cell’s cytoplasm covered by the plasma membrane is displaced outward to the myelin sheath and forms the membranous covering of the fiber, the neurolemma. Small gaps, called nodes of Ranvier, are formed between adjacent Schwann cells. – Unmyelinated axons are formed when Schwann cells do not tightly wrap axons. In unmyelinated axons, a single Schwann cell usually associates with several axons and only partially encloses the axons. Figure 17.9 Illustration showing structural differences between myelinated and unmyelinated axons in the peripheral nervous system. 5 Myelinated and Unmyelinated Axons of the CNS • In the central nervous system, the neuroglia called oligodendrocytes associate with the axons to form myelinated axons. – Oligodendrocytes associate with several axons by produce sheet-like extensions that wrap around, thus, myelinating the axons. – Mostly, myelinated fibers are organized into the areas of white matter of the brain and spinal cord. – Unmyelinated fibers are common in the gray matter of the brain and spinal cord. Figure 17.10 Illustration showing myelination of fibers (axons) in the central nervous system by oligodendrocyte. Lab Activity 2 – Medullated Nerve • Observe a microscopic preparation labeled “Medullated nerve, teased.” – A nerve is a part of the peripheral nervous system that consists of parallel axons (fibers) and their associated Schwann cells enclosed in connective tissue wrappings. – Teasing the nerve separates the axons (fibers) for individual observation. Some preparations are specifically prepared (treated with osmic acid) to show the internal details of the myelin sheath, the node of Ranvier, and the axon. Otherwise, the preparation will usually show only the surfaces of the Schwann cells and the nodes of Ranvier. Figure 17.11 Illustration of teased nerve fibers. Lab Activity 2 – Medullated Nerve • Schwann cells – Schwann cells are located sequentially along the axon. Each Schwann cell tightly wraps the axon to form a myelin sheath. The Schwann cell’s cytoplasm surrounded by plasma membrane are located to the outside of the myelin sheath and are called the neurilemma. Figure 17.12 High power photograph of teased nerve fibers (at the node of Ranvier). • Myelin sheath – The myelin sheath is formed by the wrapping of the myelin containing plasma membrane of the Schwann cell around the axon. The myelin sheath is easily observed in preparations treated with osmic acid. Myelin treated with osmic acid is darkly stained. If not treated with osmic acid, myelin sheaths are identified as “remnants.” 6 Lab Activity 2 – Medullated Nerve • Nodes of Ranvier – Nodes of Ranvier are gaps formed between adjacent Schwann cells. They allow exposure of the axon to the extracellular environment. NERVE • Axon – An axon is the process (branch) of a neuron which • (1) generates and • (2) conducts nerve impulses, and • (3) releases neurotransmitter at its terminal synapses. A nerve is a part of the peripheral nervous system and consists of parallel axons (fibers) and their associated Schwann cells enclosed in connective tissue wrappings (sheaths). – The portions of the axons in nerves are the long processes (fibers) that function in conduction of nerve impulses. Nerves • Nerves may contain Organization of a Nerve • • Each axon (fiber) and its associated Schwann cells are surrounded by a connective tissue sheath called the endoneurium. A connective tissue sheath called the perineurium organizes individual fibers (axons) and their associated endoneuria into groups called fascicles. An outermost connective tissue sheath called the epineurium organizes the fascicles into a nerve. Epineurium • Perineurium • Fascicles • Endoneurium – (1) only myelinated fibers, – (2) only unmyelinated fibers, or – (3) a combination of both. • According to the directions of impulse conduction, nerves are classified as – (1) sensory - Sensory nerves contain fibers of sensory (afferent) neurons that convey impulses to the central nervous system – (2) motor - Motor nerves contain fibers of motor (efferent) neurons and convey impulses away from the central nervous system to effectors. – (3) mixed (both sensory and motor) - Mixed nerves contain a mixture of both sensory and motor fibers. – The epineurium is the outer connective tissue sheath of the nerve. The epineurium surrounds groups of fibers (axons) called fascicles. – The perineurium is the connective tissue sheath that organizes fibers into fascicles. – Fascicles are groups of fibers surrounded by the connective tissue sheath called the perineurium. – The endoneurium is the inner connective tissue sheath that surrounds each individual axon (fiber). Organization of a Nerve • Axons – The axon functions in the generation and the conduction of the nerve impulse, and releases a neurotransmitter at its terminal synapses. The portions of the axons in nerves are the long processes (fibers) that function in conduction of nerve impulses. – All axons of the peripheral nervous system are associated with Schwann cells. • In a section of a nerve, an axon is identified as a tiny dark circular structure associated with a Schwann cell. Myelinated axons are centrally located within the myelin sheaths of the Schwann cells. Axons and their associated Schwann cells are surrounded by a connective tissue sheath called the endoneurium. Organization of a Nerve • Schwann cells – Schwann cells are associated with the axons of the peripheral nervous system. Myelinated axons are formed by the wrapping of the myelin containing plasma membranes of Schwann cells around the axons. Unmyelinated axons are formed when a Schwann cell associates with several axons and does not tightly wrap the axons. • Neurilemma – The neurilemma is the membranous covering of a nerve fiber. The neurilemma is formed by the thin region of cytoplasm and the plasma membranes of the Schwann cells. • Myelin sheath – Myelin sheaths are the fatty sheaths that are formed by the tight wrapping of the Schwann cells around the axon. Unless the specimen is specifically stained for myelin (medullated nerve with osmic acid), the myelin sheath is lightly stained and exists as a “remnant.” 7 Figure 17.13 General structure of a nerve. A nerve is a part of the peripheral nervous system and consists of parallel axons (fibers) and their associated Schwann cells enclosed in connective tissue wrappings. Figure 17.14 Illustration of a cross section of a nerve. A nerve is a part of the peripheral nervous system and consists of parallel axons (fibers) and their associated Schwann cells enclosed in connective tissue wrappings. Lab Activity 3 Cross Section of a Nerve • Observe a preparation of a “Nerve, c.s.” (c.s.- cross section). A cross section of a nerve may be presented singly on a slide preparation or may be accompanied by a longitudinal section (l.s.). Preparations with both sections are typically labeled “Nerve, c.s. & l.s.” • Most general preparation of nerves are not prepared with osmic acid, thus, do not show darkly stained myelin sheaths. Instead the myelin sheaths are observed as “remnants” and are mostly identified by location. • Observe the preparation with scanning power and identify the epineurium, perineurium, fascicles, axons, and myelin sheaths. Figure 17.16 Illustration of myelinated fibers from a cross section (c.s.) of a nerve (high magnification). The myelin sheaths of the Schwann cells are not well preserved and are identified as “remnants.” Figure 17.15 Scanning power photograph of a cross section of a nerve. Since general preparations are not treated with osmic acid, the myelin sheaths are observed as “remnants.” Figure 17.17 High power photograph of fibers (axons) from a cross section of a nerve. Preparation does not show preserved myelin sheaths. 8 Lab Activity 4 Longitudinal Section (l.s.) of a Nerve Figure 17.18 High power photograph of fibers (axons) from a cross section of a nerve prepared with “Masson” stain. Preparation does not show preserved myelin sheaths. • Observe a microscopic preparation labeled “Nerve, l.s.” A longitudinal section of a nerve is most useful in showing the relationship between the Schwann cells and the axon. • As with the cross section, myelin sheaths are observed only if the specimen was processed to maintain the myelin (medullated nerve treated with osmic acid). Otherwise, the myelin sheaths exist as “remnants” and are lightly stained. Figure 17.20 High power photograph of fibers (axons) from a longitudinal section of a nerve. Figure 17.19 Low power photograph of a longitudinal section of a nerve. Lab Activity 5 Cross Section of Medullated (Myelinated) Nerve • Observe a microscopic preparation labeled “Medullated Nerve, c.s., osmic acid” (c.s.- cross section). The observation of a nerve in cross section allows a study of – (1) its connective tissue organization and – (2) of axons. • Myelinated axons are best observed in preparations of a “medullated nerve” treated with osmic acid. Identify the – – – – – – – – (1) epineurium, (2) perineurium, (3) endoneurium (4) fascicles, (5) axons, (6) Schwann cells, (7) neurilemma, and (8) myelin sheaths. Lab Activity 5 - Myelinated Nerve (with myelin sheaths stained) • Observe a preparation of a “Medullated Nerve, osmic acid.” Treatment of medullated (myelinated) nerve preparations with osmic acid darkly stains the myelin sheaths. • Observe the preparation with scanning power and identify the – – – – – epineurium, perineurium, fascicles, axons, and myelin sheath. 9 Figure 17.21 Scanning power photograph of a cross section of a medullated nerve prepared with osmic acid. Myelin sheaths are easy to identify because osmic acid stains myelin black. Figure 17.22 Illustration of myelinated fibers from a cross section of a medullated nerve (high magnification). Schwann cells surrounding the axons have regions called myelin sheaths and regions of cytoplasm with a covering plasma membrane, the neurilemma. SPECIALIZED NEURON ENDINGS Figure 17.23 High power photograph of medullated fibers (axons) from a cross section of a medullated nerve. Most of the axons are surrounded by thick myelin sheaths. Receptors • Specialized neuron endings of the peripheral nervous system are found associated with the (1) sensory (afferent) and (2) motor (efferent) neurons. • The sensory division of the PNS relies on dendrites, the receptive portion of the axon, to respond to stimuli. – Dendrites may be modified into specialized structures called receptors. – The motor division of the PNS relies upon the synaptic transmission of the nerve impulse from the axon to the effector, a muscle or a gland. Specialized neuron endings of the peripheral nervous system are found associated with the (1) sensory (afferent) and (2) motor (efferent) neurons. Receptors • Receptors are sensory endings which respond to specific types of stimuli. – A stimulus is a change that promotes a response. For sensory receptors stimuli are mediated through a change in the receptors environment. – Among the stimuli that receptors respond to are changes in temperature (thermoreceptors), mechanical forces such as pressure and stretch (mechanoreceptors), and chemicals such as acids and electrolytes (chemoreceptors). 10 Effectors • Effectors are the muscles and glands controlled by the peripheral nervous system. – The axons of efferent neurons synapse with effectors and rely upon a neurotransmitter to mediate the flow of information (nerve impulse). – Synapses with muscles are the neuromuscular junctions and with glands the neuroglandular junctions. Figure 17.24 Simplified neural pathway between a receptor (Pacinian corpuscle) and an effector (neuromuscular junction - neuron synapse with skeletal muscle fiber) RECEPTOR Pacinian Corpuscle RECEPTOR Pacinian Corpuscle RECEPTOR Pacinian Corpuscle – Upon stimulation, the dendrite generates an electrical potential called a graded potential. – A graded potential is a local response (here restricted to the dendrite), and produces an electrical signal that has an intensity related to the strength of stimulation. If the graded potential reaches an intensity sufficient to stimulate the axon to threshold, an action potential is produced and propagated to the axon’s terminus, the axon terminals. – The axon terminals in response to the action potential release a neurotransmitter, which functions as a chemical mediator for the transfer of the electrical information. • Among the many specialized receptors of the peripheral nervous system, the Pacinian corpuscle is large and distinct. – Pacinian corpuscles are lamellated pressure receptors (mechanoreceptor) mostly located deep in the dermis of the skin and in the loose connective tissues distributed throughout the body. – A Pacinian corpuscle consists of a centrally located dendrite (the receptive portion of the neuron) surrounded by layers of flattened Schwann cells, which are surrounded by a connective tissue capsule. Lab Activity 6 Pacinian Corpuscle • A Pacinian corpuscle has a dendrite (receptive region) located in the center of concentric layers of flattened Schwann cells (lamellae), which is surrounded by a connective tissue capsule. 11 Lab Activity 6 Pacinian Corpuscle • Dendrite – The centrally located dendrite is the receptive portion of the Pacinian corpuscle. • Capsule – The capsule is the outer connective tissue layer of the Pacinian corpuscle. • Lamellae – The lamellae are concentric layers of flattened Schwann cells. Figure 17.25 A Pacinian corpuscle is a pressure receptor (mechanoreceptor). It has a dendrite (receptive region) located in the center of concentric layers of flattened Schwann cells (lamellae), which is surrounded by a connective tissue capsule. Effector Neuromuscular Junctions • EFFECTOR • Neuromuscular Junctions • The axon of a motor neuron may branch many times as it enters a muscle. At the point where an axon approaches the muscle fiber (cell), it branches into many small terminal branches (telodendria) that end in knob-like axonal terminals. The site on the muscle cell where the axon’s terminals come in close contact (synapse) with the muscle fiber is called the neuromuscular junction. A small space, called the synaptic cleft, separates the axon’s terminals and the adjacent region of the muscle fiber, the motor end plate. The motor end plate is the specialized region of the muscle fiber’s plasma membrane that contains receptors for the neurotransmitter. Thus, the neuromuscular junction includes the axon’s terminals, synaptic cleft, and the motor end plate. Lab Activity 7 Neuromuscular Junction • Observe a preparation labeled “Neuromuscular junctions.” The preparations of neuromuscular junctions are of skeletal muscle fibers (cells). Follow several axons, each to its junction with a skeletal muscle fiber. Figure 17.26 Scanning power photograph of neuromuscular junctions. A neuromuscular junction consists of the axon’s terminals, a synaptic cleft, and the motor end plate of the skeletal muscle cell (fiber) 12 Lab Activity 7 Neuromuscular Junction • Axon – The axons of motor neurons are the processes that generate and conduct nerve impulses to the neuromuscular junctions and at their axonal terminals release neurotransmitter. • Neuromuscular junction Figure 17.27 High power photograph of a neuromuscular junction. – The neuromuscular junction consists of the axon’s terminals, a synaptic cleft, and the motor end plate of the skeletal muscle cell (fiber). Electrical Terminology • Potential energy – State of electrical energy as measured by the potential to produce electrical effects • Voltage (potential) Physiology of Conduction – Electrical measurement used to describe electrical potential between two points. • Current – Flow of electrical charge and is due to the electrical difference (voltage) between two points • Resistance – Opposition to electrical flow • Insulators have high resistance • Conductors have low resistance Electrical Terminology • How might the following terms apply to these two batteries? – Potential energy • Are both the same? – Voltage • Are both the same? – Current Electrical Terminology and the Cell Membrane • How might the following terms apply to the illustrated cell membrane? – Potential energy – Voltage – Current – Resistance Extracellular • Do both produce the same? Size AAA Size D – Resistance • Does a battery contain a “resister?” Intracellular 13 Electrical Terminology and the Cell Membrane • How might the following terms apply to the illustrated cell membrane? – – – – Extracellular Potential energy Voltage Current Resistance • How might the following terms apply to the illustrated cell membrane? – – – – Intracellular Membrane Potentials Extracellular Electrical Terminology and the Cell Membrane Extracellular Potential energy Voltage Current Resistance Intracellular Resting Membrane Potential • Ionic difference between intracellular and extracellular fluids – Extracellular higher concentration of Na+ (and Cl-) – Intracellular higher concentration of K+ and negative proteins. Net result is potential difference between extracellular and intracellular. Intracellular – Extracellular is positive (Na+) – Intracellular is negative due to negative proteins. Membrane Potential Changes Mechanical Channels • If the resting membrane potential is to change – must be a change in the distribution of positive and/or negative charges; a redistribution of ions – • Movement of ions can result when ions move through channels which include • Sodium channels (typical) open when subjected to mechanical stimulus – Mechanically-gated (regulated) channels • Open when subjected to a mechanical stimulus – Voltage-gated (regulated) channels • Open when subjected to an electrical stimulus – Chemically-gated (regulated) channels • Open when subjected to a specific chemical such as a neurotransmitter or hormone – Passive (leakage) channels • Ions may leak through channels (or the phospholipid bilayer) 14 Channels • Identify regions which are – Mechanically gated – Electrically gated – Chemically gated Generator Potential • Local response (graded potential at stretch receptor) • Sodium ions move across membrane • Interior becomes less negative (more positive) • Depolarization (changes toward less negative (positive) voltage – May not reach threshold, thus no effect (action potential) – May reach threshold and produce an action potential Threshold and Action Potentials • Threshold – Point of depolarization (stimulation) which initiates an effect (action potential) – In this case the electrically-gated Na+ channels open, (which are adjacent to the active mechanically-gated channels). – The mechanically gated Na+ channels become inactive • Action potential – Not local; travels great distance – Involves electrically-gated channels – Propagated along fiber (axon) Depolarization as Na+ Moves Inward • Receptor’s Na+ channels become inactive • Local current opened adjacent electricallygated Na+ channels (threshold) • These channels produce local current Adj tN + Generation of Action Potential 1. Resting membrane potential is established 2. Depolarization phase – Increase in sodium ion permeability – Self propagating event 3. Repolarization phase – Decrease in sodium ion permeability – Increase in potassium ion permeability • • Undershoot or after-hyperpolarization occurs Redistribution of sodium and potassium by ATP driven sodium-potassium pump Repolarization as K+ Moves Out • Local current opens adjacent electrically-gated K+ channels • K+ moves outward and repolarization occurs • Local currents open adjacent Na+ channels • Action potential is propagated to adjacent forward ti 15 Na+ / K+ Pump • The Na+ / K+ reestablishes the extracellular and intracellular ionic gradients – Pump requires ATP – Na+ is pumped outward – K+ is pumped inward Synapse • Anatomical relationship between neurons, or neurons and an effector organ, and at which a nerve impulse is transmitted through the action of a neurotransmitter. Synapse Components of Synapse • Consist of – Presynaptic membrane of axonal terminal (synaptic knob or bouton) which functions in the release of neurotransmitter – Postsynaptic membrane (of dendrite, postsynaptic neuron, effector, organ, etc.) which houses receptors for neurotransmitter – Synaptic cleft of extracellular material between presynaptic and postsynaptic membranes which electrically isolates the membranes. Termination of Neurotransmitter • Enzymes associated with postsynaptic membrane or present in cleft • Reuptake by astrocytes into presynaptic terminal where degraded by enzymes • Neurotransmitter diffuses away from synapse 2. Calcium ion channels open 1. Action potential arrives 3. Calcium ions promote exocytosis of neurotransmitter, calcium ions are quickly removed 4. Neurotransmitter binds to postsynaptic receptors 5. Receptors allow passage of specific type of ions 6. Depending upon ion movement postsynaptic membrane is either 7. Neurotransmitter is deactivated depolarized (EPSP) or hyperpolarized (IPSP) by enzymatic action; some components may be reused IPSP and EPSP • EPSP • Excitatory postsynaptic potential results when interior becomes more positive • IPSP • Inhibitory post-synaptic potential results when interior becomes more negative 16 Synaps e Synaps e A B The result is “A” or “B”? Which is produced? A) action potential, B) IPSP, C) EPSP? Summation • Summation is the adding together of synaptic potentials (SPs). Could be EPSPs, IPSPs, or both EPSPs and IPSPs. • Temporal summation – Pertaining to time; the quick succession of SPs at a few synapses are summated • Spatial summation A B The result is “A” or “B”? Which is produced? A) action potential, B) IPSP, C) EPSP? Mechanisms of Neurotransmitters • Direct acting – Channel linked receptors result in the opening of ion channels – – Alter membrane potential of target – Can produce depolarization (sodium ions move inward) and hyperpolarization (potassium ions move outward) – Pertaining to space; many SPs occur over the postsynaptic membrane and are summated Mechanisms of Neurotransmitters • Indirect acting – Involves G-protein complex – Results in the production of a second messenger – Second messenger may influence enzymes to Autonomic Regulation • Activate or inactivate proteins (translation) • Regulate gene activity (transcription) • Regulate membrane ion channels and potentials 17 Autonomic Systems Brain Motor Division (efferent PNS) CNS Parasympathetic (autonomic, visceral) Sympathetic (autonomic, visceral) Spinal cord • Sympathetic – “fight or flight response” – Terminal neurotransmitter is epinephrine (E) or norepinephrine (NE) • Parasympathetic Cranial nerves Somatic (skeletal muscle, voluntary) PNS Spinal nerves Sympathetic (sweat glands, involuntary) – “resting and digesting,” or “rest and repose” – Terminal neurotransmitter is acetylcholine (ACh) • Organs – May have dual innervations, response is excitation by one system and inhibition by other system – May have single innervations, response is promoted or not promoted. HUMAN BRAIN • Four major regions of the human brain, HUMAN BRAIN Four major regions of the human brain, (1) cerebrum, (2) cerebellum, (3) diencephalon, and (4) brain stem. Figure 17.28 Midsagittal view of human brain showing four major regions. – (1) cerebrum, – (2) cerebellum, – (3) diencephalon, and – (4) brain stem. Figure 17.29 Lateral view of human brain. 18 Figure 17.30 Superior view of human brain. Figure 17.32 Illustration showing a midsagittal section of the human brain. Figure 17.31 Inferior view of human brain. Figure 17.33 Photograph of a midsagittal section of the human brain. Cerebrum • The cerebrum is the largest part of the brain. CEREBRUM The cerebrum is the largest part of the brain. – The cerebrum is divided into the right and left cerebral hemispheres, which are connected inferiorly by a large band of white matter, the corpus callosum. – The cerebrum functions include • integrating somatic (body) sensory and motor information, • thought, • memory, • reason, and • emotions. 19 Cerebrum • Cerebral hemispheres – The right and left cerebral hemispheres form the superior portion of the brain. Externally, four lobes, the (1) frontal, (2) parietal, (3) occipital, and (4) temporal, are named for both their associated cranial bones and cerebral landmarks. Both hemispheres are referred to as the cerebrum. • Gyri – Gyri are rounded elevated ridges on the surface of the cerebrum. • Sulci Figure 17.34 Functional areas (Brodmann areas) of the cerebrum. – Sulci are shallow grooves on the surface of the cerebrum. • Fissure – Fissures are deep furrows. Two dominate fissures of the brain are the longitudinal fissure and the transverse fissure. Cerebrum • Longitudinal fissure – The longitudinal fissure is a deep groove that medially divides the cerebrum into its right and left cerebral hemispheres. • Transverse fissure – The transverse fissure separates the superiorly located cerebrum from the inferiorly located cerebellum. • Central sulcus – A central sulcus is the centrally located shallow groove of each cerebral hemisphere that divides each frontal lobe from each parietal lobe. Cerebrum • Frontal lobes – A frontal lobe is the most anterior lobe of each cerebral hemisphere. Each is separated posteriorly from a parietal lobe by shallow groove, a central sulcus. – The functional regions of the frontal lobes include • (1) somatic motor cortex that controls movement of skeletal muscles, • (2) a premotor cortex for learned (memorized) motor skills and habits • (3) a motor area (Broca’s area) for motor control of muscles associated with speech, • (4) cognition (process of knowing - awareness, perception, reasoning, and judgment), • (5) language centers for word association and meaning. Cerebrum Cerebrum • Precentral gyrus – The precentral gyrus of each frontal lobe is a rounded elevated ridge located immediately anterior to each central sulcus. • A precentral gyrus functions as the primary motor cortex and houses the neurons (pyramidal cells) directly involved in conscious control of skeletal muscles. • Parietal lobes – Each parietal lobe is located posterior to its associated central sulcus and anterior to each occipital lobe. – The primary function of the parietal lobes is housing the areas that receive and integrate relayed somatic sensory information, the somatosensory areas. – The primary sensory (somatosensory) areas of the postcentral gyri receive the somatosensory information. – Posterior to each postcentral gyrus, the somatosensory association areas function to integrate the sensory information so that it is comprehensible. – The parietal lobe also functions in sensory integration for spatial visualization (visual attention) and manipulation of objects. • Postcentral gyrus – The postcentral gyrus of each parietal lobe is a rounded elevated ridge located posterior to each central sulcus. A postcentral gyrus functions as the primary sensory (somatosensory) cortex as it houses the neurons that receive information relayed from receptors in the skin and from a group of receptors distributed in muscles, tendons, and joints, the proprioceptors. • Occipital lobes – The occipital lobes are located posterior to the parietal lobes. Each occipital lobe is separated anteriorly from its associated parietal lobe mostly by the parieto-occipital fissure. The function of the occipital lobes is to house the visual cortex. The posterior occipital lobe houses the primary visual cortex, the neurons that receive information relayed from the visual fields of the eyes. Anterior to the primary visual cortex, the visual association area allows interpretation of received visual information. 20 Cerebrum • Temporal lobes Cerebrum • Cerebral cortex – Each temporal lobe is located laterally on each hemisphere. Each is separated from the frontal and parietal lobes by the lateral sulcus. The functions of the temporal lobes include the (1) auditory areas, (2) language area (Wernicke’s area), (3) memory, and the (4) ability to categorize objects. – The primary auditory cortex is located in the superior portion of each temporal lobe and receives information relayed from the auditory receptors. Inferior to the primary auditory cortex the auditory association area allows meaningful interpretation of the auditory information. – Wernicke’s area functions in recognition of spoken words. • Lateral sulcus – Each lateral sulcus separates each temporal lobe from its associated frontal and parietal lobes. – The cerebral cortex is the outer gray matter of the hemispheres. It is composed mostly of neuron cell bodies and unmyelinated fibers. • Cerebral white matter – The cerebral white matter is deep to the gray matter. It is composed mostly of myelinated nerve fibers. • Lateral ventricles – The two lateral ventricles are large chambers, one of which is located within each cerebral hemisphere. The lateral ventricles communicate with the third ventricle, each by way of an interventricular foramen. Each lateral ventricle houses the choroid plexus, which extends from the third ventricle. The choroid plexus is the site for the production and regulation of cerebrospinal fluid (CSF). Cerebrospinal fluid fills the lateral ventricles and drains into the third ventricle. Cerebrum • Corpus callosum – The corpus callosum consists of fibers (white matter) that connect corresponding areas of the right and left cerebral hemispheres. It is located superior to the lateral ventricles and deep in the longitudinal fissure. It functions in communication between the right and left cerebral hemispheres. • Fornix – The fornix is an arched band of white matter located inferior to the corpus callosum. It connects and functions in communication between regions of the brain called the hippocampus (functions in memory processes) and the hypothalamus. • Septum pellucidum – The septum pellucidum is a membrane that medially separates the two lateral ventricles. DIENCEPHALON The diencephalon is a region of the brain that is surrounded by the cerebral hemispheres. It consists mostly of the (1) thalamus, (2) hypothalamus, and (3) epithalamus. Diencephalon • Thalami – The thalami are two interconnected regions of gray matter, the right thalamus and the left thalamus, that form the superior portions of the lateral walls of the third ventricle. A bridge of fibers, the intermediate mass, passes across the third ventricle and connects the two thalamic regions. The primary function of the thalami is to relay incoming sensory information to various regions of the cerebral cortex. • Epithalamus Figure 17.39 The diencephalon consists of the epithalamus, thalami, and the hypothalamus. – The epithalamus is located superior to the thalamus and forms the thin roof of the third ventricle. The pineal gland (body) extends outward from the posterior surface of the epithalamus. • Pineal gland (body) – The pineal gland is a posterior extension of the epithalamus. The pineal gland functions as an endocrine gland releasing melatonin, a hormone that functions in sleep cycles and reproduction. 21 Diencephalon • Hypothalamus – The hypothalamus is located inferior to the thalamus. It forms the inferior portions of the lateral walls and the floor of the third ventricle. Among the functions of the hypothalamus are: • (1) regulation of body temperature, • (2) sensations (drives) of hunger and thirst, • (3) production of two hormones, antidiuretic hormone (ADH) and oxytocin (OT), that are released at the posterior pituitary gland, • (4) production of regulatory hormones to control the anterior pituitary gland, • (5) and regulation of the autonomic nervous system (especially involving the responses to stress and the coordination of information flow between the pons and medulla). Diencephalon • Infundibulum – The infundibulum is a hollow stalk of tissue which extends from the hypothalamus to the pituitary gland. The infundibulum serves as a pathway for (1) fibers (axons) and (2) blood vessels that leave the hypothalamus and enter the pituitary gland. – Fibers (axons) from specialized endocrine producing neurons of the hypothalamus pass through the infundibulum, terminate at the posterior pituitary gland, and release the hormones (1) antidiuretic hormone (ADH) and (2) oxytocin (OT). – A specialized unit of blood vessels in the hypothalamus functions to pickup regulatory hormones. The blood vessels pass through the infundibulum and deliver the regulatory hormones to the anterior pituitary. The regulatory hormones function in controlling the secretory activity of the anterior pituitary. Diencephalon • Third ventricle – The third ventricle is a narrow chamber centrally located in the diencephalon. The third ventricle houses a choroid plexus, a structure that produces cerebrospinal fluid (CSF). In addition to functioning as a site for the production of cerebrospinal fluid by its choroid plexus, the third ventricle receives cerebrospinal fluid from the lateral ventricles. From the third ventricle cerebrospinal fluid drains into the cerebral aqueduct of the midbrain. BRAIN STEM The brain stem is located between the diencephalon and the spinal cord. Superiorly to inferiorly, the brain stem consists of the (1) midbrain, (2) pons, and the (3) medulla oblongata. Brain Stem • Midbrain – Located immediately inferior to the diencephalon, the midbrain is the most superior portion of the brain stem. Functions of the midbrain include (1) serves as a pathway for ascending and descending tracts (peduncles), (2) visual and auditory reflexes (corpora quadrigemina), (3) control of muscle tone, (4) regulation of cerebral nuclei, and (5) maintenance of consciousness. Two cerebral peduncles (bundles of myelinated fibers) that contain ascending and descending fiber tracts are located on the midbrain's ventral-lateral surface. The posterior surface of the midbrain is formed by four rounded elevations called the corpora quadrigemina. The midbrain also houses several nuclei (red nucleus and substantia nigra). Figure 17.40 The brain stem consists of the midbrain, pons, and the medulla. The corpora quadrigemina is the posterior region of the midbrain. The cerebral aqueduct is a channel that delivers CSF to the fourth ventricle. 22 Brain Stem • Corpora quadrigemina Brain Stem • Pons – The corpora quadrigemina are four rounded elevations that form the posterior surface of the midbrain. – The superior two elevations are the superior colliculi and the inferior elevations are the inferior colliculi. – The pons is the rounded bulge of the brain stem located between the midbrain (superior) and the medulla oblongata (inferior). The pons functions to • (1) connect the cerebellum superiorly with the midbrain and cerebrum and inferiorly with the medulla oblongata and spinal cord, • (2) contain nuclei for respiration and • (3) contain nuclei of four cranial nerves (trigeminal- V, abducens- VI, facial- VII, and vestibulocochlear nerves- VIII). • The superior colliculi contain nuclei that mostly function in visual reflexes, especially involving movement of the head, neck, and eyes. • The inferior colliculi contain nuclei that function in auditory reflexes, especially involving the movement of the head, neck, and extremities. Brain Stem • Medulla oblongata Brain Stem • – The medulla oblongata is located between the pons (superior) and the spinal cord (inferior). Its ventral surface is composed of two ridges of motor fibers called the pyramids. The medulla oblongata functions include: – (1) connects the spinal cord with the brain (medulla is connection into the brain stem), – (2) houses autonomic centers (nuclei) that function in the regulation of respiration and circulation (cardiovascular center - control of heart rate and force of contraction and blood vessel diameter (tone), – (3) contains nuclei of five cranial nerves (vestibulocochlear-VIII, glossopharyngeal- IX, vagus- X, accessory- XI, and hypoglossal- XII nerves), – (4) relay centers for somatic sensory information to the thalamus and cerebellum, and – (5) functions as the site where motor tracts (pyramidal tracts) from the motor cortex cross to the opposite side, the decussation of the pyramids. Cerebral aqueduct – The cerebral aqueduct is narrow channel within the midbrain that connects the third and the fourth ventricle. – The cerebral aqueduct drains cerebrospinal fluid from the third ventricle to the fourth ventricle. • Fourth ventricle – The fourth ventricle is located posterior to the pons. Superiorly, the fourth ventricle is continuous with the cerebral aqueduct, and inferiorly it is continuous with the central canal of the spinal cord. – The fourth ventricle houses a choroid plexus, a structure that produces cerebrospinal fluid (CSF). – In addition to functioning as a site for the production of cerebrospinal fluid by its choroid plexus, the fourth ventricle receives cerebrospinal fluid from the third ventricle by way of the cerebral aqueduct. – Cerebrospinal fluid moves out of the fourth ventricle into the subarachnoid space, a space formed under the arachnoid meninx, a membrane covering that surrounds the brain and the spinal cord. Cerebellum Cerebellum The cerebellum is located posterior to the pons and medulla and inferior to the occipital lobes of the cerebral hemispheres. • The cerebellum is located posterior to the pons and medulla and inferior to the occipital lobes of the cerebral hemispheres. • The cerebellum functions include the coordination of complex muscle movements and the maintenance of posture and balance. 23 Cerebellum • Cerebellar hemispheres – The cerebellar hemispheres are located one on each side of the cerebellum’s central vermis. • Vermis – The cerebellar vermis is the central region of the cerebellum that functions to connect the two cerebellar hemispheres. • Folia Figure 17.41 The cerebellum functions include the coordination of complex muscle movement and the maintenance of posture and balance. – Folia are the horizontally oriented rounded ridges of cerebellar hemispheres. The folia are separated by narrow grooves called sulci. • Arbor vitae – The arbor vitae are the branching areas of cerebellar white matter. VENTRICLES The ventricles of the brain are small interconnected internal cavities that contain the choroid plexuses and cerebrospinal fluid (CSF). Figure 17.46 The ventricles of the brain. Ventricles • Lateral ventricles – A large lateral ventricle is found within each cerebral hemisphere. The two ventricles are medially separated by a thin partition called the septum pellucidum. The lateral ventricles contain a structure that functions in the production and regulation of cerebrospinal fluid (CSF), the choroid plexus. The lateral ventricles (first and second) communicate with the third ventricle each by an interventricular foramen, which allows CSF to flow into the third ventricle. • Third ventricle – The third ventricle is a narrow chamber located within the diencephalon. It receives CSF for its own choroid plexus and from the lateral ventricles. From the third ventricle CSF flows into the cerebral aqueduct to the fourth ventricle. Ventricles • Fourth ventricle – The fourth ventricle is located between the pons and the cerebellum. It connects to the cerebral aqueduct superiorly and is continuous with the central canal of the spinal cord. The fourth ventricle receives CSF from its own choroid plexus and from the cerebral aqueduct. From the fourth ventricle CSF flows through foramina into the subarachnoid space, where the CSF then circulates around the brain and spinal cord. 24 Meninges • The meninges are the three membranes that surround the brain (and spinal cord). MENINGES The meninges are the three membranes that surround the brain (and spinal cord). – The meninges protect and isolate the brain (and spinal cord) by their structure and the spaces they form, and they provide routes for blood vessels. – From outer to inner the three meninges are the • (1) dura mater, • (2) arachnoid, and • (3) pia mater Meninges • Dura mater – The dura mater is the outermost of the meninges. It is a tough fibrous membrane that is continuous around the brain and spinal cord. The outer surface of the brains dura mater is attached to the periosteum (lining of bone). The inner surface of the dura contains blood vessels and specialized veins called dural sinuses. – One of the dural sinuses, the superior sagittal sinus is located at the mid-line of the dura’s superior surface. In addition to receiving venous return, the superior sagittal sinus receives cerebrospinal fluid (CSF). A minute space, the subdural space, separates the dura mater from the underlying arachnoid. Meninges • Arachnoid – The arachnoid is the middle meninx. The subarachnoid space contains cerebrospinal fluid and the arachnoid trabeculae. The arachnoid trabeculae are a network of collagen and elastic fibers that function to support the arachnoid and the maintain the subarachnoid space. Cerebrospinal fluid functions to protect the spinal cord and allow diffusion of various chemical substances. • Pia mater – The pia mater is the innermost of the meninges. It is a delicate vascular membrane that adheres to the brain (and spinal cord). Cerebrospinal Fluid • Cerebrospinal fluid, or CSF, is produced by the choroid plexuses located in the brains ventricles. CEREBROSPINAL FLUID Cerebrospinal fluid, or CSF, is produced by the choroid plexuses located in the brains ventricles. – A choroid plexus is a vascular proliferation that produces CSF by a combination of vascular filtration and regulation by neuroglia called ependymal cells. – CSF exits the fourth ventricle by way of three foramina and enters the subarachnoid space, where it circulates around the brain and spinal cord. – Specialized areas of the arachnoid meninx, the arachnoid granulations, penetrate the dura mater of the superior sagittal sinus and allow drainage of the CSF into venous circulation. 25 Cranial Nerves • There are twelve pairs of cranial nerves, each named and numbered with a roman numeral. – Except for the first two cranial nerves, ten pairs emerge from the brain stem. Figure 17.47 Overview of circulation of cerebrospinal fluid. Sheep Brain Dissection DISSECTION OF THE SHEEP BRAIN Because of its small size, ease of storage, and most importantly, its anatomical simularites to the human brain, the preserved sheep brain is ideal for dissection Sheep Brain Dissection EXTERNAL ANATOMY • Because the sheep is a quadruped, directional terms of reference are obviously different for the sheep brain than for the human (biped). Liberty is taken and direction terms that apply to the sheep brain in biped orientation are used (for example, anterior (biped) is substituted for cranial (quadruped). EXTERNAL VIEWS Sheep brains may be purchased with all the meninges and the pituitary gland present or absent. It is important at this time to make observations to determine which meninges are present and whether or not the pituitary is present. 26 Meninges • The brain and spinal cord are surrounded by three connective tissue membranes called the meninges. • From outer to inner, they are the Dura mater • Observe the brain for the presence of the dura mater, the outer meningeal membrane which covers the brain as a tough opaque membrane. – If the membrane is present, you will not be able to directly observe the surface of the brain. (If the dura mater is not present on the preserved brain, proceed to the identification of the arachnoid). – Use scissors and carefully make an incision across the dorsal surface of the brain about an inch back from the brain’s anterior margin. – Along the midline of the brain, observe that the dura forms a large sagittal sinus (blood vessel) and forms a partition between the two cerebral hemispheres. – (1) dura mater, – (2) the arachnoid, and – (3) the pia mater. Sheep brains may be purchased with or without the dura mater (and arachnoid). All brains have the pia mater. Arachnoid • All brains should have an intact arachnoid. The arachnoid appears as a thin semitransparent layer on the surface of the brain. – Blood vessels in the sulci are observed under the arachnoid. Figure 17.49 Superior surface of the sheep brain showing the dura mater. Pia mater • Figure 17.51 Anterior surface of the sheep brain showing the arachnoid. All brains should have an intact pia mater. The pia mater is a delicate membrane which adheres to the surface of the brain. Blood vessels may be seen associated with it. The pia mater may be separated from the brain by carefully picking the surface of the brain with a probe. 27 Figure 17.53 Lateral surface of the sheep brain. Figure 17.52 Superior (dorsal) surface of the sheep brain. External Structures • Gyri External Structures • Longitudinal fissure – Gyri are elevated ridges on the surfaces of the cerebral hemispheres. • Sulci – The longitudinal fissure is a deep groove that medially divides the cerebrum into its right and left cerebral hemispheres. • Cerebellum – Sulci are shallow grooves on the surfaces of the cerebral hemispheres. • Cerebrum (cerebral hemispheres) – The right and left cerebral hemispheres form the dorsal portion of the brain. Both hemispheres are referred to as the cerebrum and are separated by the longitudinal fissure. Without tearing the two hemispheres apart, carefully separate the fissure and observe the corpus callosum. The corpus callosum connects the two cerebral hemispheres. – The cerebellum is located posterior to the cerebrum. It is separated from the cerebrum by the transverse fissure. • Transverse fissure – The transverse fissure separates the cerebrum from the cerebellum. • Spinal cord – The spinal cord is not part of the brain. It is seen extending posteriorly from the brain stem. External Structures • Pituitary gland – The sheep brain is purchased either with or without the pituitary gland. The pituitary gland is observed as a small, dome-shaped mass of tissue protruding from the ventral surface of the brain. It is located immediately posterior to the optic chiasma. (If the brain does not have a pituitary gland, proceed to the next identification). – If the brain was purchased without the complete dura mater, a small remnant of the dura is preserved around the pituitary gland. – Without removing the pituitary, gently lift its posterior aspect (along with the dura mater) away from the brain. Notice that it is attached anteriorly to the brain (hypothalamus) by a small stalk, the infundibulum. Remove the pituitary and observe the hollow infundibulum. Remove any remaining dura mater. Posterior View • Without tearing the cerebrum and the cerebellum apart, carefully bend the cerebellum away from the cerebrum to expose the – pineal gland and the – corpora quadrigemina. 28 Figure 17.54 Posterior surface of the sheep brain. Figure 17.55 Photograph of posterior surface of the sheep brain. External Structures • Corpora quadrigemina – The corpora quadrigemina are four rounded elevations on the posterior surface of the midbrain. • Pineal gland (body) – The pineal gland (body) is a rounded gland (body) that extends from the posterior border of the epithalamus. Figure 17.56 Photograph of inferior surface of the sheep brain with dura mater surrounding the pituitary gland. Ventral Views • Observe the inferior aspect of the sheep brain. Observe the brain to determine if the brain has the pituitary gland attached to its inferior surface (read previous discussion of “pituitary gland”). Figure 17.57 Photograph of inferior surface of the sheep brain with dura mater removed from the pituitary gland. 29 External Structures • Longitudinal fissure – The longitudinal fissure is a deep grove that medially divides the cerebrum into its right and left cerebral hemispheres. • Olfactory bulb – The olfactory bulbs are located on the inferior surface of the frontal lobes of the cerebrum. They receive the olfactory nerve fibers which pass through the cribriform plate of the ethmoid bone from the olfactory epithelium. • Optic nerve – The optic nerves are sensory nerves whose axons arise from the retina. • Optic chiasma – The optic chiasma is the area where the two optic nerves meet. Axons from the medial aspect of each eye cross over to the opposite side. Figure 17.58 Photograph of inferior surface of the sheep brain the pituitary gland removed. External Structures • Optic tract – The optic tract is the pathway of axons into the brain from the optic nerve and the optic chiasma. • Mamillary body – The mamillary body is a rounded body that protrudes from the ventral surface of the hypothalamus. • Infundibulum – A stalk of the hypothalamus that is located posterior to the optic chiasma and anterior to the mamillary body. It provides a connection between the hypothalamus and the posterior lobe of the pituitary gland (the pituitary gland is not shown on the illustration). External Structures • Brain Stem – The brain stem is located between the diencephalon (epithalamus, thalamus, and hypothalamus) and the spinal cord. Anteriorly to posteriorly, it consists of the midbrain, pons, and the medulla oblongata. • Midbrain – The midbrain is located posterior and slightly inferior to the diencephalon, which is centrally located between the two cerebral hemispheres. The ventral surface is composed of ascending and descending fiber tracts called cerebral peduncles. • Pons – The rounded bulge of the brain stem is located between the midbrain (anterior) and the medulla oblongata (posterior). External Structures • Medulla oblongata – The medulla oblongata is located between the pons (anterior) and the spinal cord (posterior). • Spinal cord – The spinal cord is not part of the brain. It is seen extending posteriorly from the brain stem. • Cranial nerves – There are twelve pairs of cranial nerves. Most of the nerves are small and delicate, and many were torn away from the brain during its removal. Carefully inspect the brain and identify as many cranial nerves as possible. INTERNAL ANATOMY The internal anatomy of the sheep brain is studied from a brain cut in equal right and left halves, a (1) midsagittal section, and a (2) transverse section. 30 Midsagittal Section of Sheep Brain Figure 17.60 Illustration showing structures of midsagittal section. Figure 17.59 A midsagittal section of the sheep brain showing four major regions of organization: (1) cerebrum, (2) cerebellum, (3), diencephalon, and (4) brain stem CEREBRUM and Associated Structures Figure 17.61 Photograph of dissected sheep brain showing structures of midsagittal section. Cerebrum and Associated Structures • Cerebrum (cerebral hemispheres) – The right and left cerebral hemispheres form the dorsal portion of the brain. Both hemispheres are referred to as the cerebrum. • Gyri – Gyri are rounded elevated ridges (convolutions) on the surface of the cerebrum. • Sulci – Sulci are shallow grooves (furrows) on the surface of the cerebrum. The cerebrum is the largest part of the brain. The cerebrum is divided into the right and left cerebral hemispheres, which are connected inferiorly by a large band of white matter, the corpus callosum. Cerebrum and Associated Structures • Transverse fissure – The transverse fissure is a deep furrow that separates the cerebrum and the cerebellum. • Cerebral and cerebellar cortex – The cortex is the outer gray matter of the cerebral hemispheres and the cerebellum. It is composed mostly of neuron cell bodies and unmyelinated fibers. Observe the outer gray matter of the cerebellum, the cerebellar cortex. • Cerebral and cerebellar white matter – The white matter is deep to the gray matter. It is composed mostly of myelinated nerve fibers. Observe the inner white matter of the cerebellum, the arbor vitae. • Lateral ventricles – The two lateral ventricles are large chambers, one of which is located within each cerebral hemisphere. They are filled with cerebrospinal fluid. 31 Cerebrum and Associated Structures • Corpus callosum – The corpus callosum consists of fibers (white matter) that connect corresponding areas of the right and left cerebral hemispheres. It is located superior to the lateral ventricles and deep in the longitudinal fissure. DIENCEPHALON and Associated Structures • Fornix – The fornix is an arched band of white matter located inferior to the corpus callosum. It connects a region of the brain called the hippocampus with the hypothalamus. • Septum pellucidum – The septum pellucidum is a membrane that medially separates the two lateral ventricles. The diencephalon mostly consists of the (1) epithalamus (2) thalami (pleural of thalamus), and hypothalamus. Diencephalon • Thalami – The thalami consists of two interconnected regions of gray matter, the right thalamus and the left thalamus, that form the superior portions of the lateral walls of the third ventricle. A bridge of fibers, the intermediate mass, passes across the third ventricle and connects the two thalamic regions. • Epithalamus – The epithalamus is located superior to the thalamus and forms the thin roof of the third ventricle. The pineal gland (body) extends outward from its posterior surface. • Hypothalamus – The hypothalamus is located inferior to the thalamus. It forms the inferior portions of the lateral walls and the floor of the third ventricle. BRAIN STEM and Associated Structures The brain stem is located between the diencephalon and the spinal cord. Anteriorly to posteriorly, it consists of the (1) midbrain, (2) pons, and the (3) medulla oblongata. • Third ventricle – The third ventricle is a narrow chamber centrally located in the diencephalon. Brain Stem • • The brain stem is located between the diencephalon and the spinal cord. Anteriorly to posteriorly, it consists of the (1) midbrain, (2) pons, and the (3) medulla oblongata. Midbrain – The midbrain is located immediately posterior to the diencephalon. On its anterior surface are the two cerebral peduncles which contain ascending and descending fiber tracts. On its posterior surface are four rounded elevations called the corpora quadrigemina. • Corpora quadrigemina • Pons – The corpora quadrigemina are four rounded elevations on the posterior surface of the midbrain. – The pons is the rounded bulge of the brain stem located between the midbrain (anterior) and the medulla oblongata (posterior). • Medulla oblongata – The medulla oblongata is located between the pons (anterior) and the spinal cord (posterior). Brain Stem • Cerebral aqueduct – The cerebral aqueduct is a narrow channel that connects the third and the fourth ventricles. • Pineal gland (body) – The pineal gland (body) is a rounded gland that extends from the posterior border of the epithalamus. • Fourth ventricle – The fourth ventricle is located inferior to the cerebellum. It connects to the cerebral aqueduct superiorly and is continuous with the central canal of the spinal cord. 32 Cerebellum • Cerebellum CEREBELLUM – The cerebellum is located superior to the pons and medulla and posterior to the cerebral hemispheres. • Arbor vitae – The arbor vitae are the branching areas of cerebellar white matter. Transverse Section of Sheep Brain Figure 17.62 Transverse section of the sheep brain. Transverse Section of Sheep Brain • Cerebral cortex – The cerebral cortex is the outer gray matter of the hemispheres. It is composed mostly of neuron cell bodies and unmyelinated fibers. Observe the cortex with a human brain model that has been sectioned to show the interior of the hemispheres. • Cerebral white matter – The cerebral white matter is deep to the gray matter. It is composed mostly of myelinated nerve fibers. Observe the white matter with a human brain model that has been sectioned to show the interior of the hemispheres. • Choroid plexus – A choroid plexus is the site of cerebrospinal fluid production. A choroid plexus is located in third ventricle and extends into the two lateral ventricles. Additionally, a choroid plexus is found in the fourth ventricle. A choroid plexus consists of capillaries and glial cells called ependymal cells. SPINAL CORD The spinal cord functions to receive, integrate, and transmit information to and from the peripheral nervous system, upper and lower levels of the cord itself, and the brain. 33 Spinal Cord • The spinal cord is housed within the vertebral canal of the vertebral column. – The spinal cord begins at the base of the brain stem with the termination of the medulla oblongata. – The spinal cord continues through the vertebral canal to its termination at the tip of its conus medullaris. – In the adult, the cone shaped ending of the spinal cord, the conus medullaris, is located at the level of the first-second lumbar vertebrae. – At the tip of the conus medullaris a fibrous strand, the filum terminale extends to the second sacral vertebra and functions to anchor the spinal cord. Meninges of Spinal Cord • Surrounding the spinal cord are three membranes called the meninges. The meninges protect and isolate the spinal cord (and brain) by their structure and the spaces they form. From outer to inner the three meninges are the – (1) dura mater, – (2) arachnoid, and – (3) pia mater. Spinal Cord – Along the spinal cord, structures called spinal roots merge to form the spinal nerves. – There are 31 pairs of spinal nerves with each pair (except spinal nerves designated C1, which exit between the skull and the first cervical vertebra) exiting the vertebral column between adjacent vertebrae. Thus, the spinal nerves are named by their vertebral associations forming the spinal nerves C1 C8, T1 - T12, L1- L5, and S1 - S5. (In the adult the regions of the spinal cord and associated spinal nerves don’t anatomically align with the regions of the vertebral column because during youth, the spinal column grows faster than the spinal cord.) Meninges of Spinal Cord • Dura mater – The dura mater is the outermost of the meninges. It is a tough fibrous membrane that is continuous around the spinal cord and brain. In the spinal cord the dura is separated from the surrounding vertebrae by the epidural space. – The epidural space mostly contains adipose tissue and serves as a passageway for blood vessels. The dura mater surrounding the brain is attached to the periosteum (lines cranial bones) and thus, does not present an epidural space. A minute space, the subdural space separates the dura mater from the underlying arachnoid. Meninges of Spinal Cord • Arachnoid – The arachnoid is the middle delicate meninx. The subarachnoid space contains cerebrospinal fluid (CSF) and the arachnoid trabeculae. The arachnoid trabeculae are a network of collagen and elastic fibers that function to support the arachnoid and the maintain the subarachnoid space. Cerebrospinal fluid functions to protect the spinal cord and allow diffusion of various chemical substances. • Pia mater – The pia mater is the innermost of the meninges. It is a delicate vascular membrane that adheres to the spinal cord (and brain). Figure 17.64 Illustration showing the relationship of the meninges and spinal cord to a vertebra (cervical region). 34 Spinal Cord • The inside of the cord is organized into gray matter and its outside into white matter. • The gray matter is divided into – – – – Figure 17.65 Scanning power photograph showing the relationship of the meninges and spinal cord in a vertebra. (1) the posterior horns, (2) the lateral horns, (3) the anterior horns, and (4) the gray commissure. • The white matter is divided into funiculi (columns), which are named according to their position as the – (1) posterior, – (2) lateral, and – (3) anterior funiculi. Spinal Cord • Spinal roots are located along the cord. Figure 17.66 Illustration showing the general structure of the spinal cord. – The dorsal root is located dorsally and consists of axons of sensory neurons. The dorsal root is associated with an enlarged region called the dorsal root ganglion. The dorsal root ganglion houses the cell bodies of sensory (unipolar) neurons. – The ventral root is located ventrally and consists of axons of motor neurons. – The dorsal and ventral roots unite to form the spinal nerves. A pair of spinal nerves exit the vertebral column between adjacent vertebrae (except spinal nerve, C1). – The spinal nerves are components of the peripheral nervous system Lab Activity 13 Spinal Cord Figure 17.67 Illustration showing the relationship of sensory and motor neurons to the organization of the spinal cord. • Observe a microscopic preparation labeled “Spinal cord, cross section.” Different quality microscopic preparations are available. Some are special “ultra thin” sections with special stains for advanced study, and others are for a general anatomical study. • Also, there are a variety of preparations from different locations along the cord. They range from having no spinal roots, to parts of roots, to complete roots with spinal nerves. 35 Figure 17.68 Illustration of the general structure of the spinal cord as seen in cross section through the spinal ganglia. Structures of Spinal Cord • Posterior median sulcus – The posterior median sulcus ia a posterior, medial, groove that partially divides the cord into right and left sides. It is narrower than the anterior median fissure. Often on histology slides it appears as a single line because it was closed by adherence of the sides. • Anterior median fissure – The anterior median fissure is an anterior, medial furrow that partially divides the cord into right and left sides. It is wider than the posterior median fissure. Figure 17.69 Scanning power photograph of a cross section of a spinal cord through the spinal ganglia (“Spinal cord ganglion, silver”). Structures of Spinal Cord • Dorsal roots – The dorsal roots contain axons of sensory neurons that synapse with the cell bodies of association neurons located in the posterior (dorsal) horns. The dorsal roots may not be present on your preparation. • Dorsal root ganglia – Each dorsal root has a dorsal root ganglion that consists mostly of cell bodies of sensory neurons and neuroglia. The cell bodies of these sensory neurons are unipolar and are surrounded by neuroglia cells. If a dorsal root ganglion is present, observe it under high magnification. It consists of the cell bodies of sensory (unipolar) neurons and their associated neuroglia cells (satellite cells). The axon directly associated with the cell body may have been cut away. • Ventral root – The ventral roots are the pathways for the axons that leave motor neurons located in the anterior (ventral) horns. Figure 17.70 Photograph showing the relationships of the dorsal root, dorsal root ganglion, ventral root, and spinal nerve to the spinal cord. Figure 17.71 Photograph showing the detail of the dorsal root ganglion. Inset shows the cell body of a unipolar neuron (sensory neuron). 36 Spinal Cord – Gray Matter • The gray matter is located to the inside of the cord and consists mostly of multipolar neurons (mostly association and motor neurons), unmyelinated fibers, and neuroglia. Figure 17.72 Scanning power photograph of the junction between the gray matter and the white matter. – The gray matter is often described as being in the shape of “butterfly wings” or the letter “H.” – The two posterior projections are called the posterior (dorsal) horns, and the two anterior projections are called the anterior (ventral) horns. – Lateral horns are seen in the thoracic and upper lumbar regions. Spinal Cord – Gray Matter • Anterior (ventral) horns – The anterior projections of gray matter are called the anterior (ventral) horns. They contain mostly the cell bodies of motor neurons and neuroglia. • Posterior (dorsal) horns – The posterior projections of gray matter are called the posterior (dorsal) horns. They contain mostly association neurons (interneurons) and neuroglia. • Gray commissure – The gray commissure is the gray matter that connects the right and left gray masses (horns) together. It encloses the central canal. Figure 17.73 High power photograph of the gray matter of the spinal cord. Spinal Cord – White matter • The white matter is located to the outside of the spinal cord’s gray matter. – The white matter contains mostly myelinated axons (some unmyelinated axons) and neuroglia. Some pathways are established by horizontal axons from one side of the cord to the other. – However, most pathways are formed by axons that are arranged along the longitudinal (vertical) axis of the cord. These axons establish pathways of ascending and descending tracts. Figure 17.74 High power photograph of the white matter of the spinal cord. Most of the axons are cut in cross section and show “remnants” of their myelin sheaths. 37 Spinal Cord – White matter • The tracts of the white matter are organized into three major white columns, or funiculi: the posterior, lateral, and anterior funiculi. • Posterior funiculi – The posterior funiculi (dorsal white columns) are located at the posterior aspect of the white matter and contain ascending tracts. • Lateral funiculi – The lateral funiculi (lateral white columns) are located at the lateral aspects of the white matter and contain ascending and descending tracts. • Anterior funiculi – The anterior funiculi (anterior white columns) are located at the anterior aspect of the white matter and contain ascending and descending tracts. 38